In Lectotype: PERSOONIA Published by the Rijksherbarium, Leiden Volume 2, Part 3, pp. 321-330 (1962) Studies on DiscomycetesII. On four species of Fimaria J. van Brummelen Rijks her barium, Leiden (With five Text-figures) Four species offimaria Vel. are described. The followingnew combinations are made: Fimaria hepatica (Batsch per Pers.) Brumm., F. cervaria (Phill. apud J. Stevenson) Brumm., and F. theioleuca (Roll.) Brumm. Ascobolus vinosus Berk., A. fuckelii J. Kunze, A. piceus (Limminghe) ex Cooke, and Fimaria murina Vel. are reduced to the synonymy of Fimaria hepatica. While investigating the taxonomy of the dark-spored Ascobolaceae the author came across a number of species that had been wrongly placed in the genus Ascobolus. Among these was a small group of closely related species of operculate Discomycetes with smooth, hyaline ascospores, transferred by various authors to such genera as Ascophanus, Coprobia, and Humaria sensu auct. non Fuckel (= Octospora Hedw. per S. F. Gray emend. Korf). Close examination of herbarium specimens or fresh material of four different species shows that there is only a superficial resemblance with Ascophanus and that the affinity with Coprobia and Octospora is rather remote. Fimaria Velenovsky of which the author was able to study the type is a valid generic name available for these species. FIMARIA Vel. Fimaria Vel., Monogr. Discom. Boh. i: 331. 1934. Fimaria murina Vel. Velenovsky's generic description of Fimaria runs: "Genus, omnino generi Humaria affine, sed apoth. margine late membranaceo-limbata, plerumque in cilias acutas fimbriata, paraph, simpl.-filif. vel ramosae. fimis, raro in limis." Fimaria was published with four species, two of which are also described in this paper. Although Velenovsky's generic description is rather clear, a more detailed one is given here: Apothecia superficial or partly immersed, often sessile on a narrow base, sometimes with a short stalk, up to mm 5 across. Receptacle at first closed and subglobular, then opening and expanding, with a broad membranaceous, lobed, dentate, crenulate or fimbriate margin, usually covered with a thin net-work of septate, branched hyphae, often reddish-brown, purplish-brown, brown or pale yellowish-brown. Excipulum clearly differentiated, of large cells, with intercellular pigment. Asc.i operculate, cylindrical, with short stalk, rounded above, the wall not bluein Melzer's reagent. Ascospores uniseriate, shortly ellipsoid to oblong-ellipsoid, at first colourless, finally with yellowish-brown contents, without oil drops or granules when mature, 321 321
Peziza Type: Humarina Ascophanus Ascophanusfuckelii Humaria Illustrative Aleuria Ascophanus 322 Persoonia Vol. 2, Part 3, 1962 smooth. Paraphyses very thin, filiform, hyaline, with the upper parts often surrounded by intercellular, amorphous pigment. On dung or on soil contaminated with dung. The amount of pigment in the fruit-bodies varies considerably even in the same species. The yellowish-brown contents of ripe ascospores show a strong fluorescence in ultra-violet light, a phenomenon Ascobolaceae. common to many species of Humariaceae and The genus differs from Ascophanus in the form of the asci, the structure of the excipulum, and the development of the fruit-bodies. Some species of Fimaria show a superficial resemblance with Ascophanus because of their occurrence on dung or on soil contaminated with it, and also because the hyaline tips of the ripe asci surrounded by a dark pigment give the disk a dotted or even roughened appearance (Fig. ib). Fimaria may be separated from Octospora and Coprobia chiefly by its different pigmentation, the typical development of the fruit-bodies, and the broad membranaceous margin of the receptacle. The genus should be placed in the Humariaceae near Octospora and Lamprospora. Fimaria hepatica (Batsch per Pers.) Brumm., comb. nov. Peziza hepatica Batsch, Elench. Fung. Contin. i: 199, pi. 26, f. 138. 1786 (devalidated name). hepatica Batsch per Pers., Mycol. eur. 1: 243. 1822. Pers.) Gillet, Champ, framj. 208. 1886. Fung. 8: 140. 1889. Eur. 76. 1907. 139. 1928. hepatica (Batsch per hepatica (Batsch per Pers.) Sacc., Syll. hepaticus (Batsch per Pers.) Boud., Hist. Class. Discom. hepatica (Batsch per Pers.) Seaver, North Am. cup-fungi, Operc. represented by Batsch I.e. fig. 138. specimen: Broome, Bowood, Bathford, Wiltshire, Great Britain, I. 1864, in Rabenhorst, Fungi eur. exs. No. 612 (Peziza hepatica). Ascobolus vinosus Berk, in Hooker, Engl. Fl. 5 (2): 2og. 1836. Dennis, Brit. Cup Fungi 41. i960. vinosus (Berk.) Holotype: Berkeley, "on a mole-hill overgrown with and covered with rabbits' and sheeps' dung", England (K-A1927).* moss, [Ascobolus testaceus (Moug. apud Fr.) Wallr. sensu Fuckel in Jb. nass. Ver. Naturk. 27-28: 58. 1873; Fuckel, Fungi rhen., fasc. 12, No. 2680. 1874; not Ascobolus testaceus (Moug. apud Fr.) Wallr. sensu Moug. in Wallr., Fl. Crypt. Germ. 2:513. 1833; nor Ascobolus testaceus P. Henn. in Hedwigia 41: 32. 1902. ] Ascobolus fuckelii J. Kunze, Fungi sel. fasc. 3, No. 286. 1879; in Oest. bot. Z. 30: 67. 1880. Ed. 2, Pilze 3: 1090. 1895. (J. Kunze) Rehm in Rabenh., Krypt.-Fl., Type: Fuckel, Fungi rhen. No. 2680. [Peziza picea Limminghe in Herb. ] Ascobolus piceus (Limminghe) ex Cooke 21: 74. 1893. Type: Limminghe, on sandy soil, s. loc., I. 1818 ("Peziza picea Limminghe", BM-A28i5, K-A1998). in Grevillea in herb. Fimaria murina Vel., Monogr. Discom. Boh. 1: 331; 2: pi. 24, f. 15. 1934. Lectotype: PR 150853 (as Boudiera murina). MISAPPLIED NAME.Humaria testacea (Moug. apud Fr.) J. Schroeter sensu J. Schroeter in Cohn, Krypt.-Fl. Schles. 3 (2): 36. 1893; Pilze Schlesiens No. 1753 Fimaria hepatica. = EXCLUDED. Ascobolus vinosus Berk. & Broome in Ann. Mag. nat. Hist. 15: 448, pi. 16, by a sporulating Hypho- f. = 25. 1865 Saccobolus versicolor (P. Karst.) P. Karst. overgrown mycete (K-A1939). * For a more accurate indication of herbarium specimens, especially when insufficiently labelled, the usual abbreviation of the herbarium is followed by the author's revision-number.
Van Brummelen: Discomycetes-II 323 Ascobolus vinosus Berk, sensu Broome in Rabenhorst, Fungi eur. = No. 658. 1864 Lamprospora miniata (Cr.) De Not. (BM, G, M). Ascobolus vinosus Berk, sensu Fuckel, Fungi rhen. No. 1852. 1866; in Jb. nass. Ver. Naturk. 23-24: 289. 1870; P. Syd., march. Myc. No. 2168. 1888 = Ascophanus carneus (Pers. per Pers.) Boud. Ascobolus vinosus Berk, sensu Boudier in Ann. Sci. nat. (Bot.) V, 10: 221-222, pi. 6, f. 11. = 1869 Ascobolus roseopurpurascens Rehm. This name is used for this species by most authors of floras and handbooks since 1869. Ascobolus vinosus Berk, sensu Ellis & Everhart, N. Am. Fungi No. 2620. 1891 = Ascobolus amoenus Oudemans. Apothecia densely crowded or gregarious, superficial or partly immersed, 1 3 (-4) mm diameter, 0.3-0.8 mm high. Receptacle at first closed and subglobular, then opening by a laciniate orifice and lenticular or hemispherical, finally often less more or saucer-shaped; vinaceous-brown, pale or dark reddish-brown; smooth, often covered with a thin layer ofcolourless hyphae which are also connected with the substrate; with a rather broad, lobed or crenulate, membranaceous, often paler coloured margin which may disappear in old fruit-bodies. Disk concave, then flat, of about the same colour as the receptacle, dotted with the hyaline tips of ripe Fig. 1. Fimaria hepatica. a. Diagrammatic section of apothecium (x 40). b. Top view of hymenium (x 125). ce. Ascospores (x 1600); c. lateral view; d, e. optical section (ad from Schoeter, Pilze Schlesiens 1753, e from type of Fimaria murina). and almost ripe asci. Hymenium 170-190 ft thick. Hypothecium about 15 /u thick, but often not clearly differentiated, of closely compacted isodiametric cells 4-9 ft diameter. Flesh of varying thickness, of subglobular or oblong cells 10-26 (35) X 8-10 (-30) ft, hyaline. Excipulum 29-55 F thick, often of only a few layers of subglobular, thick-walled cells 20-35 diameter /< (textura globulosa), with intercellular, amorphous, vinaceous-brown pigment, covered with hyaline, branched hyphae 5-8 /( thick. Asci broadly cylindrical, with short a stalk, rounded above, 170-200 X 19-25 iu, 8-spored, the wall not blue in Melzer's reagent. Ascospores uniseriate, ellipsoid or oblong-ellipsoid, at first colourless, but at maturity with pale yellowish-brown contents, 22-35 (38.5) X 10-13 smooth. n, Paraphyses simple or branched, septate, cylindrical, sometimes slightly swollen underneath the septa, 2.5-3.5 /< thick, clavate at the 4-8 /< thick tip, with hyaline contents, sometimes the walls in the upper part pale brownish, covered in the upper part (up to 60 ft from the tip) by intercellular, amorphous, vinaceous-brown pigment.
1Peziza Fimaria Ascobolus Humaria 324 Persoonia Vol. 2, Part 3, 1962 HABITAT.On dung of mouse and rabbit and soil which has been in on contact with dung of these animals. ILLUSTRATIONS.Batsch, Elench. Fung. Contin. I: pi. 26, f. 138. 1786; Berkeley, Outl. Brit. Fungol. pi. 23, f. 1. i860; Berkeley & Broome in Ann. Mag. nat. Hist. ILL, 15: pi. 15, f. 19. 1865; Cooke, Mycographia pi. f. 22, 85. 1876; Patouillard, Tab. anal. Fung. 1: f. 485. 1886; Velenovsky, Monogr. Discom. Boh. 2: pi. 24, f. 15. 1934; Dennis, Brit. Cup Fungi pi. VIII, f. I. i960. SPECIMENS EXAMINED. GREAT BRITAIN: Berkeley, s. loc., s. dat., in British fungi 159 ( Ascobolus vinosus, BM, E, K); Berkeley No. 42, s. loc., s. dat. (PC-A 2312); Berkeley, Badminton, Gloucestershire, XII. 1864 (K-A1937); Berkeley, s. loc., s. dat. (K-A 1925); Berkeley, "on a mole-hill with overgrown moss, and covered with rabbits' and sheeps' dung", s. loc., s. dat._(holotype of Ascobolus vinosus, K-A 1927). BELGIUM: Libert, Malmedy, s. dat, in Reliquia Libertianae No. 95 (BR-A336). FRANCE: Leveille, on rabbit dung, Bois de Boulogne, 1850 (PC-A2315 1 ); Per soon, near Versailles, s. dat. (Ascobolus purpureus, an unpublished name, L 910.256-878). GERMANY: J. Kunze, in fields, "Oberfeld" near Eisleben, X. 1878, in Fungi sel. No. 286 (Ascobolus fuckelii, BM, BRSL, GRO). POLAND: Schroeter, on rabbit dung, Wroclaw: Oswitz, s. dat., in Pilze Schlesiens No. 1753 ( Humaria testacea, HBG). CZECHOSLOVAKIA: Velenovsky, on soil near nest of mice, Solopisky, Bohemia, 31. X. 1925 (lectotype of Fimaria murina, PR 150853); Velenovsky, on dung of mouse, Hubfchov, Mnichovice, Bohemia, X. 1927 (PR 151039). UNKNOWN COUNTRY: Limminghe, on sandy soil, s. loc., I. 1818 ("Peziza picea in herb. Limminghe", type of Ascobolus piceus, BM-A2815, K-A1998). This is probably the most common species of the genus. According to Seaver (1928: 139) this species is also found in Colorado, U.S.A. The ascospores show a considerable variation in length, sometimes even in the same fruit-body. Although Batsch's description is rather short and only based on full-grown fruit-bodies, a striking unanimity as to the conception ofhis species exists in literature. The description and illustration of Berkeley & Broome (1865: 445) certainly established the stability we find in the works of Cooke (1876: 42), Phillips (1887: 99), Rehm (1894: 946) and Seaver (1928: 139). FIMARIA LEPORUM (Alb. & Schw. per Pers.) Vel. sensu Fuckel Peziza granulosa var. ββ, leporum Alb. & Schw., Consp. Fung. 337. 1805 (devalidated name). granulosa var. leporum Alb. & Schw. per Pers., Mycol. eur. 1: 298. 1822. Pezzia leporum (Alb. & Schw. per Pers.) Fuck., Fungi rhen. 1877. 1866. - leporum (Alb. & Schw. per Pers.) Fuck, in Jb. nass. Ver. Naturk. 23-24: 288. 1870. leporum (Alb. & Schw. per Pers.) Sacc., Syll. Fung. 8: 138. 1889. Coprobia leporum (Alb. & Schw. per Pers.) Boud., Hist. Class. Discom. Eur. 69. 1907. leporum (Alb. & Schw. per Pers.) Vel., Monogr. Discom. Boh. 1: 331. 1934. Type locality: Germany, Ober Lausitz. The modern conception of this species is based on Fuckel, Fungi rhen. No. 1877. 1866; in Jb. nass. Ver. Naturk. 2324: 288. 1870. Apothecia gregarious, sessile or substipitate, 1-3 mm diameter. Receptacle at first closed and subglobular, then opening and more or less scutellate; reddishbrown or purplish-brown; almost smooth, covered with a thin net-work of hyphae, often with small fragments of the substratum adhering, with a prominent membranaceous margin almost disappearing with age. Disk concave or flat, brown or
a. Van Brummelen: Discomycetes-H 325 Ascophanus Humaria purplish-brown, smooth, dotted with the hyaline tips of ripe asci. Hymenium about 140 ft thick. Hypothecium about 15 ft thick, of closely compacted hyphae 1.7-3.5 P thick, also with some larger cells up to 13 X 7.5 Flesh of varying thickness, ft. up to 210 ft thick in the centre consisting of strongly branched hyphae with oblong or irregularly shaped cells 10-24 x 5 -I 4 t* ( f which the contents strongly stain in cotton blue or congo red) and of subglobular cells 10-30 ft diameter, hyaline. Excipulum 15-30 ft thick, of rather thick-walled, subglobular or oblong cells 9 -I 5 x 5 _I 3 1" (textura globulosa or textura angularis), brownish, with intercellular pigment and pigment in the cell-walls, covered with a net-work of strongly branched hyphae 3-5.5 ft thick. Asci cylindrical, with a short stalk, rounded above, up to about 140 ft long and 14-15 ft wide, 8-spored, the wall not blue in Melzer's reagent. Ascospores uniseriate, broadly ellipsoid, at first hyaline, finally with pale yellowish-brown contents, 14.4-15.9 X 9.8-10.8 ft, smooth; sometimeswith a single, central oil drop when immature. Paraphyses branched, septate, filiform, 2.0-2.5 f 1 thick, shortly clavate and 3-5 ft thick at the tip, hyaline, with reddish- or purplishbrown, amorphous pigment among the upper parts. HABITAT.On dung of rabbit and hare. ILLUSTRATIONS.Cooke, Mycographia pi. 19, f. 75. 1875; Cooke in Grevillea 3: pi. 43, f. 201. 1875. SPECIMENS EXAMINED. J. Kunze, on rabbit dung, Nonnenthal near Eisleben, Germany, end IV. 1875, in J. Kunze, Fungi sel. No. 188 (GRO, HBG, PAD). The material in Fuckel's Fungi rhen. No. 1877 is in very poor condition. Fig. 2. Fimaria leporum. Diagrammatic section ofapothecium (x 40). bd. Ascospores (x 1600); b, c. lateral view; d. optical section. From J. Kunze, Fungi sel. No. 188 (GRO). This species is well-characterized by its broadly ellipsoid ascospores. It was also collected by Svrcek (1959: 96) from different localities in Bohemia. Fimaria cervaria (Phill. apud J. Stevenson) Brumm., comb. nov. Peziza cervaria Phill. apud J. Stevenson, Mycologia scotica 308. 1879. cervaria (Phill. apudj. Stevenson) Sacc., Syll. Fung. 8: 143. 1889. cervarius (Phill. apud J. Stevenson) Boud., Hist. Class. Discom. Eur. 76. 1907. Type locality: East Scotland, Great Britain. Apothecia gregarious, superficial, sessile or mm substipitate, 1.5-3 diameter, about 1 mm high. Receptacle at first subglobular, then less more or scutellate, with a short stalk-like base; chestnut-brown or purplish-brown; smooth and covered with a net-work ofbrown hyphae; margin thin, crenulate or fimbriate.disk concave, chestnut-brown or purplish-brown, almost smooth. Hymenium about 150 ft thick. Hypothecium about 25 ft thick, of closely compacted cells 4-10 X 3-7 ft. Flesh up to 900 ft thick in the central part, of small isodiametric cells 7-14 ft diameter and of large oblong cells 19-53 x 5-12 Pi hyaline. Excipulum near the margin
326 Persoonia Vol. 2, Part 3, 1962 25-4 I thick, 1 near the base up to 75 /i thick, of subglobular, angular or slightly elongated cells 8-40 n diameter (textura angularis or textura globulosa), with intercellular, amorphous, purplish-brown pigment, especially in the outer few layers of cells and between the hyphae which cover the excipulum; covering hyphae 2-3.5 /" thick, near the margin often forming a more or less fimbriate layer. Asci Fig. 3. Fimaria cervaria. a. Diagrammatic section of apothecium (x 40). bd. Ascospores (x 1600); b. lateralview; c, d. optical section. e. Section of margin of apothecium (x 400) (from Gremmen 1712).
Van Brummelen: Discomycetes-II 327 cylindrical, with a very short stalk, rounded above, 150-175 X 10-12.5 n, 8-spored, the wall not blue in Melzer's reagent. Ascospores uniseriate, ellipsoid or oblongellipsoid, first colourless, finally often with pale yellowish-brown contents, at 14.4-17.0 X 7.0-8.5 n, without oildrops or granules when mature, smooth. Paraphyses branched, especially in the upper parts, septate, filiform, about 2.0 /it thick, not or only very slightly enlarged, 2.0-3.0 /t thick at the tip, hyaline but near the tip often with pale brownish walls, with intercellular, amorphous, purplish-brown pigment covering the tips. HABITAT.Only known from dung of deer. ILLUSTRATIONS.None published thus far. SPECIMEN EXAMINED.GREAT BRITAIN: Gremmen 1712, on dung of deer, Loch-an- Eilean, near Aviemore, Scotland, 29. V. i960 (L, Herb. Gremmen). U.S.A.: Cain, on deer dung, Ringwood, near Ithaca, New York, 6. IX. 1952 (TRTC 24287). Both collections fully agree with Phillips' description. ac. Fig. Fimaria theioleuca. 4. Diagrammatic sections of apothecia (x 25). d. Surface view of portion of excipulum (x 750). eg. Ascospores (x 1600); e. lateral view; f, g. optical (a, b, dg from van Brummelen 648, c. from Webster s collection). section
328 Persoonia Vol. 2, Part 3, 1962 van Brum- Fig. Fimaria 5. theioleuca. Section of margin of apothecium (x 600) melen 648). (from
(i860). Van Brummelen: Discomycetes-Il 329 Fimaria theioleuca (Roll.) Brumm., comb. nov. Pseudombrophila theioleuca Rolland in Bull. Soc. mycol. Fr. 4: 57, pi. 15, f. 3a-c. 1888. - Humaria theioleuca (Roll.) Sacc., Syll. Fung. 8: 126. 1889.Type locality: France, Paris, Botanic Garden. Apothecia solitary or gregarious, substipitate or stipitate, 14 mm diameter, up to 1.5 mm high. Receptacle at first closed and subglobular, then opening and turbinate with a narrow or stalk-like base; pale yellowish-brown or darker brown; smooth or folded, covered with a net-work of branched hyphae; margin entire, membranaceous, brown, often somewhat fimbriate at the outside. Disk flat, then convex, pale yellowish, smooth. Hymenium 155175 thick. /< Hypothecium 20-30 /< thick, of closely compacted, isodiametric cells 5-9 diameter. Flesh of /«varying thickness, in the central part up to about 1000 p, of subglobular of oblong cells 11-43 X 9-28 fi, hyaline. Excipulum near the margin 25-35 <" thick, in the margin 20-28 n thick, in the lower parts up to 72 /< thick, of subglobular, thick-walled cells 11-30 (-40) diameter (textura globulosa), a with brownish cell-walls and intercellular pigment especially in the outer layers, covered with a net-work of septate, branched, hyaline or encrusted 3-9 /i thick hyphae which is often more dense and fimbriate toward the margin. Asci cylindrical, with a short stalk, rounded above, 155-180 X r 3 1 5 /I, 8-spored, the wall not blue in Melzer's reagent. Ascospores uniseriate, ellipsoid, at first colourless, finally with pale yellowish-brown contents, 13.2-15.7 X 7.7-8.3 /(, smooth, without oil drops or granules. Paraphyses branched, septate, filiform 1.21.8 fi thick, not enlarged upward, hyaline, with granular contents, without any pigment. HABITAT.Known from dung of deer, rabbit and sheep. ILLUSTRATIONS. Rolland, I.e. SPECIMENS EXAMINED.GREAT BRITAIN: on rabbit Webster, dung, University of Sheffield, 5. XII. 1961 (L). NETHERLANDS: van Brummelen 648, on dung of deer, Elspeet, Gelderland, 3. IV. '959 (L). This species was incorrectly placed by Rolland and Boudier (1907: 65) in the genus Pseudombrophila Boud. which is characterized by small clusters of short, coloured, septate, obtuse hairs on the outside of the receptacle. These hairs are quite different from the hyphae covering the outer surface of the apothecia in the present species. The collection mentioned above fully agrees with Rolland's description and certainly belongs to the genus Fimaria. LITERATURE BERKELEY, M. J. (1836). The English Flora of Sir James Edward Smith. Vol. V, Part II, Fungi. London. Outlines of British Fungology. London. BERKELEY, M.J. & BROOME, C. E. (1865). Notices of British Fungi. In Ann. Mag. nat. Hist. Ill, 15: 444-452, pi. 15-17- BOUDIER, J. L. E. (1907). Histoire et classification des Discomycetes d'europe. Paris COOKE, M. C. (1875-1879). Mycographia seu Icones Fungorum. London. DENNIS, R. W. G. (i960). British Cup Fungi and their Allies. London. FUCKEL, K. W. G. L. (1870). Symbolae mycologicae. Beitrage zur Kenntniss der rheinischen Pilze. In Jb. nass. Ver. Naturk. 23-24: 1-459, ph 1-6. PHILLIPS, W. (1887). A Manual of the British Discomycetes. London.
330 Persoonia Vol. 2, Part 3, 1962 REHM, H. (1887-1896). Ascomyceten. In Rabenh., Krypt.-Fl., Ed. 2, 1 (3). ROLLAND, L. (1888). Trois nouvelles especes de Discomycetes. In Bull. Soc. mycol. Fr. 4: 56-58, pi. 15. SEAVER, F. J. (1928). The North American Cup-fungi (Operculates). New York. STEVENSON, J. (1879). Mycologia scotica. The Fungi Distribution. Edinburgh. of Scotland and their geographical SVRCEK, M. (1959). Nekolik zajimavych druju koprofilnich hub pozorovanych v. roce 1958. In Ceska Mykologie 13: 92-102. VELENOVSKY, J. (1934). Monographia Discomycetum Bohemiae (2 parts). Praha.